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BURLINGTON, Vt. (AP) - The fatal beating of a man at a Vermont homeless encampment stemmed from a dispute between separate camps that was triggered when the victim poured bodily fluids onto the tent of one of his assailants and not because he was transgender, police said Friday.

The four people facing second degree murder charges in the May 28 death of Amos Beede were arrested late Thursday on a San Diego beach known as a destination for transients following a nationwide manhunt.

Burlington police identified the suspects, all in their 20s, as Erik Averill, Myia Barber, Allison Gee and Jordan Paul on Thursday afternoon.

At a Friday morning news conference, Burlington Police Chief Brandon del Pozo said the investigation found that a resident of one encampment smeared feces on a tent in the other encampment about 100 feet away, and Beede was assaulted after he poured urine onto Averill’s tent.

“We believe it fits into a wider patter of aggression against fellow homeless on the part of Mr. Averill rather than targeted aggression motivated by Mr. Beede’s transgender status,” del Pozo said.

But the circumstances and brutality of the attack led investigators to question at the outset whether Beede being transgender played a role. The investigation is continuing, and nothing has been ruled out.

“When somebody is beaten to death with brute force, without using weapons, just using hands and fists, which we believe at this point is what happened here, that’s a crime that takes a lot of intensity, a lot of anger, rage, commitment on the part of the perpetrators,” the chief said. “And when a person is a victim and is transgender we wonder if the motive for that type of assault is bias.”

Police said Beede had a home in Milton, but would occasionally stay at the homeless encampment not far from the city’s downtown.

The suspects fled Vermont the day after the assault. They came into contact with police in Roswell, New Mexico, after Averill was charged with assaulting another member of his group, but he was released before police realized they were wanted in Vermont.

Burlington police said Paul had connections in San Diego and police in that city located them on a beach that is a known destination of itinerants.

San Diego police Sgt. Michael Tansey told The Burlington Free Press the group will be held until authorities in Burlington bring charges against them and seek their extradition from California. They have yet to be assigned attorneys who can comment.

Beede’s family posted a statement on the website of the lesbian, gay, bisexual, transgender and queer advocacy group Pride Center of Vermont that said contrary to earlier reports he was not homeless. The statement said Beede’s being transgender was an important part of his life. It said he was a “friend to many” who went to the homeless camp to support others in need.